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Corresponding author: Daubian Santos ( daubians@gmail.com ) Academic editor: Bradley Sinclair
© 2022 Daubian Santos, Rodrigo dos Reis Santos, Guilherme Cunha Ribeiro.
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The subgenus Amphineurus (Ramphoneurus Alexander) (Diptera: Limoniidae), a group of craneflies endemic to southern South America, is revised. The previously described species are redescribed and illustrated, along with fifteen new species: A. (R.) alexanderi sp. nov., A. (R.) amorimi sp. nov., A. (R.) anchoralis sp. nov., A. (R.) anfractus sp. nov., A. (R.) billinghami sp. nov., A. (R.) caleuchus sp. nov., A. (R.) deceptus sp. nov., A. (R.) falcatus sp. nov., A. (R.) immaculatus sp. nov., A. (R.) morphyi sp. nov., A. (R.) oosterbroeki sp. nov., A. (R.) podenasi sp. nov., A. (R.) theischingeri sp. nov., A. (R.) triangularis sp. nov., and A. (R.) stigmaticus sp. nov. In addition, A. (R.) fuscifusus Alexander is considered a nomen dubium. A phylogenetic analysis with a broad taxonomic sampling with all valid species of the subgenus A. (Rhamphoneurus Alexander) and species from all the subgenera of Amphineurus Skuse is performed. The broad outgroup sampling used in the study (including taxa from the entire range of distribution of Amphineurus) suggests that the subgenus A. (Rhamphoneurus) is a monophyletic taxon, that its closest relatives are taxa currently distributed in New Zealand, and that Australasian taxa are paraphyletic with respect to the Neotropical Clade.
Chioneinae, Chile, crane flies, phylogeny, taxonomy, Tipulomorpha
The southernmost parts of the Southern Hemisphere contain a rich and unique biota of craneflies (Diptera: Tipuloidea) which is poorly understood, but that nevertheless suggests strong biogeographical links between southern South America and Australasia. In a comprehensive review of the subject,
Amphineurus was erected by
The genus Amphineurus includes four subgenera and over 80 species. Amphineurus (Amphineurus), with approximately 42 described species, is distributed in southern South America, New Zealand, New Caledonia, Australia and New Guinea. The Australian species of A. (Amphineurus) were revised by
It is of historical interest that, when reporting for the first time the occurrence of Amphineurus in the Neotropical Region,
As discussed by
In the present contribution, we provide a detailed taxonomic revision and phylogenetic study of the Neotropical subgenus A. (Rhamphoneurus), with the description of 15 new species and an identification key for all its included taxa. Knowledge of the previously described species of the subgenus A. (Rhamphoneurus) is virtually limited to the original descriptions. In addition, knowledge of the morphology and biology of immature stages of this subgenus remains unknown.
Testing the monophyly of A. (Rhamphoneurus) demanded broad taxonomic sampling including representatives of all subgenera of Amphineurus from the entire distribution range of the genus; i.e., from South America, New Zealand, Australia, Tasmania, New Guinea and New Caledonia.
The specimens of this study belong to the Alexander Collection of Crane Flies of the National Museum of Natural History, Smithsonian Institution, USA (
Descriptive terminology follows
Abbreviations used in the figures are as follows: a ej, ejaculatory apodeme; ad spur, anterodorsal spur; aed, aedeagus; aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; ap, gonocoxite apodeme; aprn, anteropronotum; arc, arculus; b spur, basal spur; cer, cercus; cgonst, clasper of gonostylus; cv, cervical sclerites; cx, coxa; db clasper, dorsal branch of clasper of gonostylus; db g, dorsal branch of gonocoxite; db lobe, dorsal branch of lobe of gonostylus; f, foramen of male tergite X; goncx, gonocoxite; gonst, gonostylus; hlt, halter; hph, hypopharynx; hyp val, hypogynial valve; ib, interbase; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lb, lobule of lobe of gonostylus; lb c, lateral branch of clasper of gonostylus; lbl, labella; lbr, labrum; lgonst, lobe of gonostylus; la spur, lateroapical spur; lb sh, lateral branch of sheath of aedeagus; lh, lateral humps of male tergite IX; mb c, medial branch of clasper of gonostylus; mh, medial humps of male tergite IX; mr, meron; ms lobe, mesal lobe of gonocoxite; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pb sh, posterior branch of sheath of aedeagus; plp, palpus; pm, paramere; pprn, postpronotum; presct, prescutum (including presutural scutum); r ms, right mesal lobe of gonocoxite; rst, rostrum; sct, postsutural scutum; sctl, scutellum; st, sternite; t r, tip of rostrum; tg, tergite; t9, male tergite IX; t10, male tergite X; vb clasper, ventral branch of clasper of gonostylus; vb g, ventral branch of gonocoxite; vb lobe, ventral branch of lobe of gonostylus; vt, vertex.
When necessary, clarifications of male and female genitalia were made with the method proposed by
The phylogenetic analysis was made under different criteria: 1) parsimony with equal weights; 2) parsimony with implied weights; 3) Bayesian inference. This was made in order to test the stability of the results under different analytical protocols.
For the parsimony analysis, we used heuristic searches with tree-bisection reconnection (TBR) branch swapping with 1,000 replicates holding up to 100 trees per replication. All characters were treated as unordered. Cases with the character state unknown were scored with ‘?’ and cases where a character is inapplicable with ‘–’. The analyses were made using TNT (
For Bayesian inference, we used the MK model (
Family Limoniidae
Subfamily Chioneinae
Amphineurus Skuse, 1890: 800 (as a subgenus of Rhypholophus). Type-species: Amphineurus (A.) umbraticus Skuse, 1890 (as Rhypholophus).
Amphineurus can be diagnosed by the following combination of characters: gonocoxite bifid; wing with R2+3+4 aligned with R2+3. When there is an element R4+5 (and therefore, no R2+3), members of the genus are distinguished from other Chioneinae by the closed discal cell.
There is great diversity in the male terminalia of Amphineurus (Figs
Schematic illustration of the male terminalia of representatives of Amphineurus (dorsal view). A A. (A.) patya; B A. (A.) superbus; C A. (A.) pita; D A. (A.) umbraticus. — Abbreviations: ad spur, anterodorsal spur; aed, aedeagus; aed sh, sheath of aedeagus; b spur, basal spur; cgonst, clasper of gonostylus; db clasper, dorsal branch of clasper of gonostylus; db lobe, dorsal branch of lobe of gonostylus; f, foramen of male tergite X; goncx, gonocoxite; la spur, latero-apical spur; lb sh, lateral branch of sheath of aedeagus; lgonst, lobe of gonostylus; lh, lateral humps; mh, medial humps; pb sh, posterior branches of sheath of aedeagus; t9, male tergite IX; t10, male tergite X; vb clasper, ventral branch of clasper of gonostylus; vb lobe, ventral branch of lobe of gonostylus.
Schematic illustration of the male terminalia of representatives of Amphineurus (dorsal view). A A. (A.) maculosus; B A. (Nothormosia) edentulus; C A. (A.) perdecorus; D A. (Nesormosia) fatuus. — Abbreviations: aed, aedeagus; aed sh, sheath of aedeagus; cgonst, clasper of gonostylus; db clasper, dorsal branch of clasper of gonostylus; goncx, gonocoxite; gonst, gonostylus; lb sh, lateral branch of sheath of aedeagus; lgonst, lobe of gonostylus; ms lobe, mesal lobe of gonocoxite; t9, male tergite IX; vb clasper, ventral branch of clasper of gonostylus.
Amphineurus has an 180º rotation of male terminalia (in the words of Edwards: “undergone permanent torsion”).
The Chioneinae genera with inverted terminalia bear great diversity and frequently they have hairy wings similar to Amphineurus. The great amount of wing vestiture resembles Maietta Alexander. The wings of some A. (Amphineurus) (Fig.
A. (Rhamphoneurus) Alexander, 1929a: 186. Type-species: A. (R.) nothofagetorum Alexander, 1929a.
This subgenus is distinguished by the rostrum at least as long as the remainder of the head. Cell M2 open.
Habitus of representative species of the subgenus A. (Rhamphoneurus) is shown in Fig.
Morphology of the head of Amphineurus (Rhamphoneurus) (male). A Head of A. (R.) chiloeanus (lateral view); B Mouthparts of A. (R.) nullus (dorsal view). — Abbreviations: cv, cervical sclerites; hph, hypopharynx; lbl, labella; lbr, labrum; plp, palpus; rst, rostrum; t r, tip rostrum; vt, vertex.
Male terminalia of Amphineurus (Rhamphoneurus) insanus. Dorsal view with some structures in lateral view. — Abbreviations: a ej, ejeculatory apodeme; aed, aedeagus; aed sh, sheath of aedeagus; ap, gonocoxite apodeme; cgonst, clasper of gonostylus; db g, dorsal branch of gonocoxite; goncx, gonocoxite; ib, interbase; l ms, left mesal lobe of gonocoxite; lb, lobule of lobe of gonostylus; lb c, lateral branch of clasper of gonostylus; lgonst, lobe of gonostylus; mb c, medial branch of clasper of gonostylus; pm, paramere; r ms, right mesal lobe of gonocoxite; vb g, ventral branch of gonocoxite.
Holotype: ♂, Chile, Llanquihue, Carelmapu [41°44′S 73°40′W], Coast of Canal de Chacao, 18-III-1957, Peña (
The species is named after C.P. Alexander, the remarkable Tipulomorpha expert who described all the species of this subgenus, among many others.
This species is distinguished by a curved R2+3, large markings only near R2, r-m longer than basal deflection of R5, and R2+3+4 longer than R2+3. Furthermore, the species has characteristic male terminalia with a trapezoidal-shaped spur and symmetrical tip of left mesal lobe.
Wing length 5.68 mm, width 1.61 mm. — Coloration: General coloration dark brown. Rostrum and palpus dark brown. First segment of antenna yellowish-brown, remaining segments dark brown. Head dark brown. Abdomen brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) alexanderi sp. nov. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E Male tergite IX (dorsal view) and remainder of male terminalia (lateral view). — Abbreviations: aed sh, sheath of aedeagus; cgonst, clasper of gonostylus; goncx, gonocoxite; l ms, left mesal lobe of gonocoxite; lgonst, lobe of gonostylus; r ms, right mesal lobe of gonocoxite; t9, male tergite IX.
Specimens of this new species were previously identified by C.P. Alexander as A. (R.) insanus Alexander. This species resembles A. (R.) insanus but differs mainly in the attachment of m-cu, length of r-m, shape of thorax, mesal lobe and sheath of aedeagus.
Holotype: ♂, Chile, Chiloé Is., Dalcahue [42°22′S 73°39′W], 10/12-II-1954, Peña (
The species is named after Dalton de Souza Amorim, a remarkable Brazilian Diptera expert.
This species is distinguished by only a marking near R2 and R2+3+4 as long as R2+3. Furthermore, the species has characteristic male terminalia with a triangular spur and an atrophied tip of the left mesal lobe.
Wing length 5.60 mm, width 1.68 mm. — Coloration: General coloration dark brown. Thorax dark brown, pleura brown. Scutum without stripes. Halter dark brown with knob pale. Coxae light brown and legs brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) amorimi sp. nov. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view). — Abbreviations: Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; t9, male tergite IX.
Specimens of this new species were previously identified by C.P. Alexander as A. (R.) insanus Alexander. This species resembles A. (R.) morphyi sp. nov. but differs mainly in the chaetotaxy of the thorax and gonocoxite, length of R2+3+4 and shape of the mesal lobe.
Holotype: ♂, Chile, Arauco, Nahuelbuta, Contulmo, Palo Botado [37°39′S 72°54′W], 12-II-1952, Peña (
The word anchoralis is Latin for “of the anchor”. The term refers to the shape of the projection of the male 9th tergite, which resembles an anchor.
The species is distinguished by the absence of markings near R1, and R2+3 shorter than R2. Furthermore, this species has a long vertical spur on the sheath of aedeagus, male tergite IX has a distinctive ridge projection, mesal lobes absent and the posterior margin of the male tergite IX is mostly unpigmented.
Wing length 6.15 mm, width 1.86 mm. — Coloration: General coloration yellowish brown. Rostrum and palpus dark brown. First segments of antenna pale, remaining segments dark yellow. Head brownish-black. Eyes black. Thorax yellowish-brown, pleura pale. Scutum without stripes. Halter pale with knobs light orange. Coxae and legs brownish. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) anchoralis sp. nov. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; k, knob of the halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; t9, male tergite IX.
Specimens of this new species were previously identified by C.P. Alexander as A. (R.) sanus Alexander and A. (R.) insanus Alexander. This species resembles A. (R.) sanus but differs mainly in M1+2, shape of projection in male tergite IX, ventral branch of gonocoxite and spur of sheath of aedeagus.
Holotype: ♂, Chile, Arauco, Nahuelbuta, Butamalal [37°49′S 73°14′W], 1100–1400 m, 23/31-I-1954, Peña (
The word anfractus in Latin is for “winding”. The term refers to the winding or sinuous aspect of the anal vein and crossvein m-cu.
This species is distinguished by sinuose crossvein m-cu and anal vein. Furthermore, this species is characterized by the male terminalia with a folded medial branch of the clasper of gonostylus and a cylindrical left mesal lobe.
Wing length 7.84 mm, width 2.48 mm. — Coloration: General coloration dark brown. Thorax dark brown, pleura brown. Scutum with one black stripe. Halter pale with knobs ochreous. Coxae brownish yellow and legs dark brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) anfractus sp. nov. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; t9, male tergite IX.
Some specimens of this new species were previously identified by C.P. Alexander as A. (R.) glabristylatus Alexander and some as A. (R.) nothofagetorum Alexander. This species resembles A. (R.) chiloeanus but differs mainly in the form of the anal vein, shape of male tergite IX, sheath of aedeagus and mesal lobe.
Amphineurus (Rhamphoneurus) apiculatus Alexander, 1968: 29 (description, fig. 15: male terminalia).
Holotype: ♂, Chile, Chaihuin [39°56′S 73°34′W], 810 m, 20-III-1955, Peña, Holotype 12206 (
This species is distinguished by having vein bM forking into M2+3+4 and M1, absence of markings on R2 and insertion of r-m desclerotized. Furthermore, the species has distinctive male terminalia with a curved sheath of aedeagus and both mesal lobes similar in length with vertically directed tiny tips.
Wing length 6.26 mm, width 1.82 mm. — Coloration: General coloration yellowish brown. Rostrum and palpus brown. First segment of antenna yellow, remainder segments brown. Head brownish gray. Eyes black. Thorax yellowish-brown, pleura brown. Scutum without stripes. Halter yellow. Coxae light yellow and legs dark yellow. Abdomen yellowish-brown. — Head (Fig.
Amphineurus (Rhamphoneurus) apiculatus Alexander. A head (dorsal view); B detail of wing; C wing; D thorax (lateral view); E thorax (dorsal view); F male tergite IX (dorsal view) and remainder of male terminalia (lateral view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; t9, male tergite IX.
This species resembles A. (R.) nonnullus Alexander but differs mainly in the fork of bM, shape of male tergite IX, clasper of gonostylus and mesal lobe.
Holotype: ♂, Chile, Osorno, Pucotrihue [40°32′S 73°42′W], 10/12-IV-1968, Peña (
The species is named after Zacariah D. Billingham, a great expert of Australian craneflies.
This species is characterized by crossvein m-cu uniformly inserted at the fork of bM. Furthermore, this species is characteristized by the male terminalia with a curved sheath of aedeagus, a small right mesal lobe and the most setose ventral branch of gonocoxite. The thorax is also distinguishable by a narrow knob of the halter and katepisternum longer than the meron.
Wing length 6.45 mm, width 1.66 mm. — Coloration: General coloration dark brown. Rostrum and palpus dark brown. First segments of the antenna yellow, remainder dark yellow. Head dark brown. Eyes blackish gray. Thorax dark brown, pleura pale. Scutum without stripes. Halter pale with knob dark yellow. Coxae brownish yellow and legs yellowish-brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) billinghami sp. nov. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; t9, male tergite IX.
Remarks. Specimens of this new species were previously identified by C.P. Alexander as A. (R.) nullus Alexander. This species resembles A. (R.) theischingeri sp. nov. but differs mainly in attachment of m-cu, setae concentration in gonocoxite, shape of male tergite IX, and mesal lobe.
Holotype: ♂, Chile, Chiloé Is. Ancud [41°52′S 73°48′W], 23-I-1932, Peña (
The word caleuchus refers to the Chilean myth of Caleuche, a sentient ghost ship of the Chiloé islands (
The species is distinguished by a darkened distal quarter of the wing, R2+3+4 similar to R2+3, and M1+2 shorter than the basal deflection of M1. Furthermore, the species is characterized by the distinctive clasper of gonostylus with setae on medial branch, curved lateral branch and male tergite IX with a very shallow notch.
Wing length 6.50 mm, width 2.05 mm. — Coloration: General coloration dark brown. Rostrum and palpus dark brown. First segments of antenna yellow, remainding segments brownish-yellow. Head brown. Eyes black. Thorax yellowish-brown, pleura yellowish brown. Scutum with three dark brown stripes. Halter pale with knob ochreous. Coxae yellow and legs brown. Abdomen dark brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) caleuchus sp. nov. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view); H female terminalia (dorsal view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cer, cercus; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; hyp val, hypogynial valve; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; ms lobe, mesal lobe of gonocoxite; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; st, sternite; tg, tergite; t9, male tergite IX.
Specimens of this new species were previously identified by C.P. Alexander as A. (R.) nothofagetorum Alexander. This species resembles A. (R.) stigmaticus sp. nov. but differs mainly in markings on the sheath of aedeagus, length of R2+3+4, and shape of lobe of gonostylus, cercus and mediotergite.
Amphineurus (Rhamphoneurus) chiloeanus Alexander, 1969: 374 (description, fig. 37: wing, fig. 38: male terminalia).
Holotype: ♂, Chile, Chiloé Is., Dalcahue [42°22′S 73°39′W], 1/4-IV-1954, Peña, Holotype 12553 (
The species is distinguished by having m-cu strongly sinuous, straight anal vein and beginning of M1+2 retains sclerotized section. Furthermore, this species has a characteristic sheath of aedeagus with a long basal spur, mesal lobe is very reduced and male tergite IX is remarkable by its rounded posterolateral lobes.
Wing length 7.37 mm, width 2.14 mm. — Coloration: General coloration light brown. Rostrum and palpus dark brown. First segment of antenna light yellow, remaining segments dark yellow. Head dark brown. Eyes black. Thorax light brown, pleura dark yellow. Scutum with three brown stripes. Halter brownish yellow with knob ochreous. Coxae brownish yellow and legs yellowish-brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) chiloeanus Alexander. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; t9, male tergite IX.
Some specimens of this species were misidentified by C.P. Alexander as A. (R.) nothofagetorum Alexander. This species resembles A. (R.) glabristylatus Alexander but differs mainly in attachment of m-cu, length of cercus, shape of male tergite IX, sheath of aedeagus and mesal lobe.
Amphineurus (Rhamphoneurus) extraordinarius Alexander, 1939: 280 (description); Alexander, 1947 (fig. 18: wing).
The species Amphineurus extraordinarius was originally included in the subgenus Amphineurus. In all the phylogenetic analysis we performed (see below in this paper), this species is the sister-group of the other species of Rhamphoneurus, and we think it should be transferred to this subgenus. This new subgenus placement is therefore based on the phylogenetic analysis, but specimens of A. extraordinarius were not available when the analysis was finalized. Consequently, a redescription of the species is not possible at this time.
Holotype: ♂, Chile, Chiloé Is. Ancud [41°52′S 73°48′W], 23-I-1952, Peña (
The word deceptus is Latin for “deceived”. The term refers to the fact that specimens of this new species were misidentified by Alexander as two different species.
This species is distinguished by the darkened distal quarter of the wing, and bM is uniform. Furthermore, the species is characterized by the distinctive male terminalia with clasper of gonostylus with setae on medial branch and straightened base of sheath of aedeagus.
Wing length 6.50 mm, width 2.06 mm. — Coloration: General coloration dark brown. Rostrum and palpus brownish-black. First segments of the antenna yellow, remainding segments dark yellow. Head dark brown. Eyes gray. Thorax dark brown, pleura yellowish brown. Scutum with three dark brown stripes. Halter brownish-yellow. Coxae brownish yellow and legs yellowish-brown. Abdomen dark brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) deceptus sp. nov. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view); H female terminalia (dorsal view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cer, cercus; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; hyp val, hypogynial valve; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; st, sternite; tg, tergite; t9, male tergite IX.
Some specimens of this new species were previously identified by C.P. Alexander as A. (R.) glabristylatus Alexander and some as A. (R.) nothofagetorum Alexander. This species resembles A. (R.) nothofagetorum but differs mainly in the shape of the branches of the clasper, male tergite IX, mesal lobes and sheath of aedeagus.
Holotype: ♂, Chile, Osorno, Bahia Mansa [40°35′S 73°44′W], South of Pucotrihue, 12-IV-1968, Peña (
The word falcatus is Latin for “armed with scythes”. This name is in reference to the sheath of aedeagus, which resembles a scythe.
This species is distinguished by absence of markings near R2 and R2+3+4 longer than R2+3. Furthermore, this species has distinctive male terminalia with a curved sheath of aedeagus, only the right mesal lobe folded, and the longest lobule of the subgenus, as long as the remainder of the lobe of gonostylus.
Wing length 6.08 mm, width 1.86 mm. — Coloration: General coloration blackish brown. Thorax blackish brown, pleura gray. Scutum without stripes. Halter ochreous. Coxae brownish yellow and legs dark brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) falcatus sp. nov. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; t9, male tergite IX.
Specimens of this new species were previously identified by C.P. Alexander as A. (R.) nullus Alexander. This species resembles A. (R.) nullus but differs mainly in the attachment of m-cu, shape of male tergite IX, prescutum and left mesal lobe.
Amphineurus (Rhamphoneurus) fuscifusus Alexander, 1929a: 190 (description).
Amphineurus (R.) fuscifusus Alexander is the only species of the genus in which the male is unknown. Additionally, the only identified specimen is the female holotype, currently housed in the Natural History Museum (
Some morphological data described by
Amphineurus (Rhamphoneurus) glabristylatus Alexander, 1929a: 188 (description, fig. 224: male terminalia).
Paratypes: 1 ♂, Chile, Chiloé Is., Mechuque Is. [42°18′S 73°16′W], 23-XII-1926, F. & M. Edwards (
This species is distinguished by the sinuous anal vein and straight m-cu. Furthermore, this species has characteristic male terminalia with a folded medial branch of the clasper of gonostylus without setae, an unpigmented male tergite IX, sinuous sheath of aedeagus, and the right mesal lobe is longer than the left mesal lobe, with a cylindrical apical tip.
Wing length 7.07 mm, width 2.22 mm. — Coloration: General coloration brown. Rostrum and palpus dark brown. First segments of antenna light yellow, remainding segments dark yellow. Head dark brown. Eyes gray. Thorax brown, pleura yellowish brown. Three reddish-brown stripes on scutum. Halter dark yellow with knob ochreous. Coxae yellowish-brown and legs brown. Abdomen brown. — Head (ventral view Fig.
Amphineurus (Rhamphoneurus) glabristylatus Alexander. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view); H female terminalia (dorsal view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cer, cercus; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; hyp val, hypogynial valve; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; st, sternite; tg, tergite; t9, male tergite IX.
We could not examine the holotype, but the paratypes were examined. This species resembles A. (R.) chiloeanus Alexander but differs mainly in the attachment of m-cu, length of female cercus, shape of male tergite IX, sheath of aedeagus and mesal lobe.
Holotype: ♂, Chile, Osorno, Pucotrihue [40°32′S 73°42 W], 10/12-IV-1968, Peña (
The word immaculatus is Latin for “unstained”. The term refers to the fact that the wing has no pterostigma or markings.
This species is characterized by having sinuous cross-vein m-cu and no markings near R2. Furthermore, the species is characterized by male terminalia with a curved sheath of aedeagus and right mesal lobe similar in shape, but longer than left one, and scutellum rounded in lateral view.
Wing length 6.22 mm, width 1.74 mm. — Coloration: General coloration brownish-yellow. Rostrum and palpus dark brown. First segments of antenna yellow, remainding segments dark yellow. Head dark brown. Eye black. Thorax brownish yellow, pleura pale. Scutum without stripes. Halter brownish-yellow. Coxae brownish yellow and legs brown. Abdomen blackish brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) immaculatus sp. nov. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view); H female terminalia (dorsal view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cer, cercus; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; hyp val, hypogynial valve; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; ms lobe, mesal lobe of gonocoxite; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; st, sternite; tg, tergite; t9, male tergite IX.
Some specimens of this new species were previously identified by C.P. Alexander as A. (R.) sanus Alexander, some as A. (R.) nullus Alexander and some as A. (R.) nonnullus Alexander. This species resembles A. (R.) billighami sp. nov. but differs mainly in attachment of m-cu, setae concentration in gonocoxite, shape of male tergite IX and right mesal lobe.
Amphineurus (Rhamphoneurus) insanus Alexander, 1952: 111 (description, fig. 9: male terminalia).
Holotype: ♂, Chile, Malleco, Rio Blanco [38°23′S 71°48′W], 5/27-III-1951, Peña, Holotype 9362 (
This species is distinguished by only a dark spot near R2 and a curved m-cu. Furthermore, this species has a remarkable thorax which is as long as tall and characteristic male terminalia with the sheath of aedeagus with a rectangular spur and the tip of left mesal lobe curved.
Wing length 6.75 mm, width 2.04 mm. — Coloration: General coloration brown. Rostrum and palpus brown. First segment of antenna yellow, remaining segments dark yellow. Head brown. Eyes black. Thorax brown, pleura yellowish brown. Scutum with one black stripe. Halter pale with knob ochreous. Coxae and legs yellowish-brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) insanus Alexander. A head (dorsal view); B detail of wing; C wing; D thorax (lateral view); E thorax (dorsal view); F male tergite IX (dorsal view) and remainder of male terminalia (lateral view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; t9, male tergite IX.
This species resembles A. (R.) alexanderi sp. nov. but differs mainly in attachment of m-cu, length of r-m, shape of thorax, mesal lobe and sheath of aedeagus.
Holotype: ♂, Chile, Chiloé Is., Dalcahue [42°22′S 73°39′W], 1/4-IV-1968, Peña (
The species is named after Charles Morphy D. dos Santos, a Brazilian entomologist who studied the taxonomy and biogeography of South American Diptera.
This species is distinguished by only a marking near R2, and basal deflection of R5 shorter than half-length of R2+3+4. Furthermore, the species is characterized by male terminalia with a triangular spur, tip of left mesal lobe bent, tip of right mesal lobe slightly curved and cercus thin and abruptly curved.
Wing length 5.63 mm, width 1.74 mm. — Coloration: General coloration yellowish brown. Rostrum and palpus dark brown. First segments of antenna yellow, remainding segments dark yellow. Head blackish brown. Eyes black. Thorax yellowish-brown, pleura yellow. Scutum with one reddish-brown stripe. Halter dark yellow with knob pale. Coxae and legs brownish yellow. Abdomen dark brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) morphyi sp. nov. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view); H female terminalia (dorsal view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cer, cercus; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; hyp val, hypogynial valve; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; ms lobe, mesal lobe of gonocoxite; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; st, sternite; tg, tergite; t9, male tergite IX.
Specimens of this new species were previously identified by C.P. Alexander as A. (R.) insanus Alexander. This species resembles A. (R.) amorimi sp. nov. but differs mainly in chaetotaxy of the thorax and gonocoxite, length of R2+3+4 and shape of mesal lobe.
Amphineurus (Rhamphoneurus) nonnullus Alexander, 1967: 491 (description); Alexander, 1968: fig. 17 (male terminalia).
Holotype: ♂, Chile, Chiloé Is., Aucar [42°9′S 73°30′W], 6/15-I-1952, Peña, Holotype 9619 (
This species is distinguished by R2 without a dark spot and the beginning of M1 desclerotized. Furthermore, the species has a distinctive male terminalia with a curved sheath of aedeagus with the branches of the clasper of different sizes.
Wing length 5.58 mm, width 1.71 mm. — Coloration: General coloration brownish-yellow. Rostrum and palpus dark brown. First segment of antenna yellow, remaining segments brownish-yellow. Head brown. Eyes grayish black. Thorax brownish yellow, pleura grayish-yellow. Scutum with one brownish-black stripe. Halter light yellow. Coxae light yellow; legs yellowish-brown. Abdomen yellowish brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) nonnullus Alexander. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; t9, male tergite IX.
This species resembles A. (R.) apiculatus Alexander but differs mainly in the fork of bM, shape of male tergite IX, clasper of gonostylus and mesal lobe.
Amphineurus (Rhamphoneurus) nothofagetorum Alexander, 1929a: 187 (description, fig. 90: wing, fig. 223: male terminalia).
Paratypes: 1 ♂, Argentina, Nahuel Huapi, Lago Correntoso [40°42′S 71°39′W], 2-XI-1926, R.C. Shannon (
The species is distinguished by the darkened distal quarter of the wings and M1+2 longer than half the length of the basal deflection of M1. Furthermore, the species has a distinctive clasper of gonostylus with setae on medial branch and straightened lateral branch.
Wing length 6.10 mm, width 1.92 mm. — Coloration: General coloration brown. Rostrum and palpus dark brown. First segments of antenna brownish yellow, remaining segments brown. Head dark brown. Eyes black. Thorax reddish-brown, pleura brown. Three brown stripes on scutum. Halter yellowish brown. Coxae and legs brownish yellow. Abdomen brown. — Head (ventral view Fig.
Amphineurus (Rhamphoneurus) nothofagetorum Alexander. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view); H female terminalia (dorsal view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cer, cercus; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; hyp val, hypogynial valve; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; st, sternite; tg, tergite; t9, male tergite IX.
We could not observe the holotype, but the paratypes were examined. This species resembles A. (R.) deceptus sp. nov. but differs mainly in shape of branches of the clasper, male tergite IX, mesal lobes and the sheath of the aedeagus.
Amphineurus (Rhamphoneurus) nullus Alexander, 1967: 492 (description); Alexander, 1968: fig. 11 (wing), fig. 16 (male terminalia).
Holotype: ♂, Chile, Chiloé Is., Aulen, Duhatao [41°58′S 74°02′W], 6/8-II-1952, Peña, Holotype 9620 (
The species is distinguished by absence of markings near R2, cross-vein m-cu attached to M3+4 and length of r-m twice length of basal deflection of R5. Furthermore, the species has distinctive male terminalia with a curved sheath of aedeagus, both mesal lobes with folded tips and branches of clasper of gonostylus of similar sizes. The species further distinguished also by the combination of sheath of aedeagus with small markings and male tergite IX pigmented. Furthermore, females of this species have the thickest cercus of the subgenus.
Wing length 6.32 mm, width 1.76 mm. — Coloration: General coloration brownish-yellow. Rostrum and palpus dark brown. First segment of antenna yellow, remainding segments dark yellow. Head grayish brown. Eyes black. Thorax brownish yellow, pleura brownish-yellow. Scutum without stripes. Halter yellow. Coxae and legs yellow. Abdomen brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) nullus Alexander. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view); H female terminalia (dorsal view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cer, cercus; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; hyp val, hypogynial valve; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; st, sternite; tg, tergite; t9, male tergite IX.
This species resembles A. (R.) falcatus sp. nov. but differs mainly in the attachment of m-cu, shape of male tergite IX, prescutum and left mesal lobe.
Holotype: ♂, Chile, Chiloé Is., Chaiten [42°58′S 72°32′W], 5/8-II-1954, Peña (
The species is named after Pjotr Oosterbroek, an important Tipulomorpha expert.
This species is distinguished by only a dark spot on R2, basal deflection of R5 as long as r-m, and M1+2 longer than basal deflection of M1. Furthermore, this species is characterized by male terminalia with sheath of aedeagus with a triangular spur and the tip of sheath inclined.
Wing length 5.47 mm, width 1.68 mm. — Coloration: General coloration brown. Rostrum and palpus brown. First segments of antenna yellow, remainding segments yellowish brown. Head brown. Abdomen yellowish brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) oosterbroeki sp. nov. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E Male tergite IX (dorsal view) and remainder of male terminalia (lateral view). — Abbreviations: aed sh, sheath of aedeagus; cgonst, clasper of gonostylus; goncx, gonocoxite; lgonst, lobe of gonostylus; ms lobe, mesal lobe of gonocoxite; t9, male tergite IX.
Specimens of this new species were previously identified by C.P. Alexander as A. (R.) insanus Alexander. This species resembles A. (R.) podenasi sp. nov. but differs mainly in the length of r-m, concentration of setae on gonocoxite, shape of male tergite IX and right mesal lobe.
Holotype. ♂, Chile, Arauco, Nahuelbuta, Pichinahuel [37°48′S 73°02′W], 1400–1600 m, 12/20-II-1953, Peña (
The species is named after Sigitas Podenas, a skilled Tipulomorpha expert.
This species is distinguished by only a dark spot on R2 and basal deflection of R5 longer than r-m. Furthermore, this species is characterized by male terminalia with sheath of aedeagus with a spur, tip of right mesal lobe curved downwards and sheath of aedeagus with rounded spur.
Wing length 5.73 mm, width 1.83 mm. — Coloration: General coloration brownish-yellow. Rostrum and palpus brown. First segment of antenna yellow, remainding segmens dark yellow. Head dark brown. Eyes black. Thorax brownish yellow, pleura pale. Contrasting golden setae on prescutum. Scutum without stripes. Halter brownish-yellow. Coxae and legs brownish yellow. Abdomen dark brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) podenasi sp. nov. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view); H female terminalia (dorsal view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cer, cercus; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; hyp val, hypogynial valve; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; ms lobe, mesal lobe of gonocoxite; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; st, sternite; tg, tergite; t9, male tergite IX.
Specimens of this new species were previously identified by C.P. Alexander as A. (R.) insanus Alexander. This species resembles A. (R.) oosterbroeki sp. nov. but differs mainly in the length of r-m, concentration of setae on the gonocoxite, shape of male tergite IX and right mesal lobe.
Amphineurus (Rhamphoneurus) rutristylus Alexander, 1968: 30 (description, fig. 18: male terminalia).
Holotype: ♂, Chile, Valdivia, Enco [39°53′S 72°8′W], 1400–1600 m, 6-III-1955, Peña, Holotype 12226 (
This species is distinguished by a dark spot near R2 and m-cu inserted at fork of bM. Furthermore this species is distinguished by the male terminalia with the lobule with two rounded margins forming heart shape, sheath of aedeagus is straight but without spurs and the mesal lobes are symmetrical, with slender, bare and slightly curved tips.
Wing length 6.87 mm, width 2.00 mm. — Coloration: General coloration dark brown. Rostrum and palpus dark brown. First segment of antenna light yellow, remainding segments dark yellow. Head brown. Eyes black. Thorax yellowish-brown, pleura light brown. Scutum without stripes. Halter brownish-yellow. Coxae brownish yellow and legs dark brown. Abdomen dark brown. — Head (Fig.
Amphineurus (Rhamphoneurus) rutristylus Alexander. A head (dorsal view); B detail of wing; C wing; D thorax (lateral view); E thorax (dorsal view); F male tergite IX (dorsal view) and remainder of male terminalia (lateral view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; t9, male tergite IX.
Some specimens of this species were misidentified by C.P. Alexander as A. (R.) insanus Alexander. This species resembles A. (R.) glabristylatus Alexander but differs mainly in the attachment of m-cu, wing markings, shape of male tergite IX and mesal lobe.
Amphineurus (Rhamphoneurus) sanus Alexander, 1929a: 189 (description, fig. 90: wing, fig. 225: male terminalia).
Holotype: 1 ♂, Chile, Chiloé Is., Ancud [41°52′S 73°48′W], 1927, F. & M. Edwards (
This species is distinguished by a subtle spot on R2. Furthermore, this species has a remarkable sheath of aedeagus with a long horizontal spur, male tergite IX with a distinctive rectangular projection, mesal lobes absent and posterior margin of male tergite IX pigmented.
Wing length 5.86 mm, width 1.90 mm. — Coloration: General coloration dark brown. Rostrum and palpus dark brown. Antenna yellow, with apical segments slightly darker. Head brown. Eyes black. Thorax dark brown, pleura brown. Scutum without stripes. Halter brownish-yellow. Coxae brownish yellow and legs dark brown. Abdomen dark brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) sanus Alexander. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; lgonst; lobe of gonostylus; mr, meron; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; sct, scutum; sctl, scutellum; t9, male tergite IX.
Some specimens were misidentified by C.P. Alexander as A. (R.) nullus Alexander. The rostrum of the holotype is longer than in the other species of this subgenus. This species resembles A. (R.) anchoralis sp. nov. but differs mainly in M1+2, shape of projection on male tergite IX, ventral branch of gonocoxite and spur of sheath of aedeagus.
Holotype: ♂, Chile, Llanquihue, Hornohuinca [41°25′S 72°37′W], XII-1968, Peña (
The word stigmaticus is Latin for “marked”. The term refers to the darkened mark near the tip of the wing of this new species.
This species is distinguished by the darkened distal quarter of the wings and basal deflection of M1 shorter than M1+2. Furthermore, the species is characterized by the male terminalia with setae on the medial branch of clasper of gonostylus and small marking on sheath of aedeagus.
Wing length 7.48 mm, width 2.57 mm. — Coloration: General coloration dark brown. Rostrum and palpus dark brown. First segment of the antenna yellow, remainding segments dark yellow. Head dark brown. Eye gray. Thorax dark brown, pleura brown. Scutum with three brownish black stripes. Halter pale. Coxae and legs brownish yellow. Abdomen dark brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) stigamticus sp. nov. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view); H female terminalia (dorsal view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cer, cercus; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; hyp val, hypogynial valve; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; ms lobe, mesal lobe of gonocoxite; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; st, sternite; tg, tergite; t9, male tergite IX.
Specimens of this new species were previously identified by C.P. Alexander as A. (R.) nothofagetorum Alexander. This species resembles A. (R.) caleuchus sp. nov. but differs mainly in marking in the sheath of aedeagus, length of R2+3+4, shape of lobe of gonostylus, cercus and mediotergite.
Holotype: ♂, Chile, Osorno, Pucotrihue [40°32′S 73°42′W], 10/12-IV-1968, Peña (
The species is named after Gunther Theischinger, an expert on Tipulomorpha who revised the Australian Amphineurus.
This species is distinguished by an absence of markings near R2, r-m as long as basal deflection of R5, and cross-vein m-cu straight and inserted at bM. Furthermore, this species is characterized by male terminalia with curved sheath of aedeagus, straight medial branch of clasper of gonostylus and mesal lobes with slender, vertical tips. The thorax is also distinguishable by narrow knob of halter and katepisternum as long as meron.
Wing length 6.30 mm, width 1.77 mm. — Coloration: General coloration dark brown. Rostrum and palpus brownish black. First segment of antenna yellow, remainding segments dark yellow. Head brownish gray. Eye black. Thorax dark-brown, pleura light brown. Scutum without stripes. Halter brownish-yellow. Coxae brownish-yellow, legs dark brown. Abdomen brownish black. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) theischingeri sp. nov. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view); H female terminalia (dorsal view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cer, cercus; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; hyp val, hypogynial valve; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; ms lobe, mesal lobe of gonocoxite; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; st, sternite; tg, tergite; t9, male tergite IX.
Specimens of this new species were previously identified by C.P. Alexander as A. (R.) nullus Alexander. This species resembles A. (R.) billighami sp. nov. but differs mainly in the attachment of m-cu, setae concentration in gonocoxite, shape of male tergite IX and mesal lobe.
Holotype: ♂, Chile, Chiloé Is., Chepu [42°2′S 74°1′W], 8-II-1952, Peña (
The word triangularis is Latin for “triangular”. This name references the triangular shape of the spur of sheath of aedeagus.
This species is distinguished by only a dark spot near R2 and M1+2 as long as the basal deflection of M1. Furthermore, this species is characterized by male terminalia with sheath of aedeagus with a triangular spur, tip of both mesal lobes bent horizontally, straightened lateral branch of clasper of gonostylus and cercus uniformly wide and abruptly angled dorsad.
Wing length 5.76 mm, width 1.74 mm. — Coloration: General coloration brownish yellow. Rostrum and palpus dark brown. First segment of antenna yellowish brown, remainding segments brown. Head brown. Eye black. Thorax brownish yellow, pleura pale. Scutum with one black stripe. Halter dark yellow with knob yellowish brown. Coxae yellow and legs brown. Abdomen yellowish brown. — Head (dorsal view Fig.
Amphineurus (Rhamphoneurus) triangularis sp. nov. A head (ventral view); B head (dorsal view); C detail of wing; D wing; E thorax (lateral view); F thorax (dorsal view); G male tergite IX (dorsal view) and remainder of male terminalia (lateral view); H female terminalia (dorsal view). — Abbreviations: aed sh, sheath of aedeagus; anatg, anatergite; anepm, anepimeron; anepst, anepisternum; aprn, anteropronotum; cer, cercus; cgonst, clasper of gonostylus; cx, coxa; goncx, gonocoxite; hlt, halter; hyp val, hypogynial valve; k, knob of halter; kepm, katepimeron; kepst, katepisternum; ktg, katatergite; l ms, left mesal lobe of gonocoxite; lgonst; lobe of gonostylus; mr, meron; mtanepst, metanepisternum; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; p, posterior basalare; patg, paratergite; pprn, postpronotum; presct, prescutum; r ms, right mesal lobe of gonocoxite; sct, scutum; sctl, scutellum; st, sternite; tg, tergite; t9, male tergite IX.
Some specimens of this new species were previously identified by C.P. Alexander as A. (R.) insanus Alexander and some as A. (R.) glabristylatus Alexander. This species resembles A. (R.) podenasi sp. nov. but differs mainly in the length of M1+2 and R2+3+4, shape of cercus and ventral branch of gonocoxite.
1 | Dark spot on R2 (Fig. |
2 |
1’ |
Faint cloudings or without markings on R2 (Fig. |
18 |
2 |
Distal quarter of wing clear (Fig. |
3 |
2’ | Distal quarter of wing darkened (Fig. |
15 |
3 |
No markings at end of R1 (Fig. |
4 |
3’ |
Dark spot distal to end of R1 (Fig. |
13 |
4 |
Antenna hairy (Fig. |
5 |
4’ |
Antenna with few setae (Fig. |
7 |
5 |
Mesal lobe of gonocoxite well-developed (Fig. |
A. (R.) rutristylus Alexander |
5’ |
Mesal lobe of gonocoxite absent (Fig. |
6 |
6 |
Spur of sheath of aedeagus projecting horizontally (Fig. |
A. (R.) sanus Alexander |
6’ |
Spur of sheath of aedeagus projecting vertically (Fig. |
A. (R.) anchoralis sp. nov. |
7 | M1+2 shorter than basal deflection of M1 | 8 |
7’ | M1+2 longer than or equal to basal deflection of M1 | 11 |
8 |
8 Tip of left mesal lobe of gonocoxite symmetrical (Fig. |
A. (R.) alexanderi sp. nov. |
8’ |
Tip of left mesal lobe of gonocoxite asymmetrical (Fig. |
9 |
9 |
Thorax as long as tall (Fig. |
A. (R.) insanus Alexander |
9’ |
Thorax longer than tall (Fig. |
10 |
10 |
Distal portion of left mesal lobe well-developed (Fig. |
A. (R.) morphyi sp. nov. |
10’ |
Distal portion of left mesal lobe atrophied (Fig. |
A. (R.) amorimi sp. nov. |
11 | R2+3+4 longer than R2+3; M1+2 as long as basal deflection of M1 | A. (R.) triangularis sp. nov. |
11’ | R2+3+4 shorter than R2+3; M1+2 longer than basal deflection of M1 | 12 |
12 | Tip of right mesal lobe projecting horizontally (Fig. |
A. (R.) oosterbroeki sp. nov. |
12’ |
Tip of right mesal lobe curved down (Fig. |
A. (R.) podenasi sp. nov. |
13 |
R2+3+4 shorter than R2+3; male tergite IX unpigmented (Fig. |
A. (R.) glabristylatus Alexander |
13’ |
R2+3+4 shorter than R2+3; male tergite IX dark (Fig. |
14 |
14 |
Anal vein straight (Fig. |
A. (R.) chiloeanus Alexander |
14’ |
Anal vein sinuose (Fig. |
A. (R.) anfractus Alexander |
15 |
Length of R2+3+4 similar to R2+3; mediotergite as wide as scutellum (Fig. |
16 |
15’ |
R2+3+4 shorter than half-length of R2+3; mediotergite narrower than scutellum (Fig. |
17 |
16 | M1+2 longer than half-length of basal deflection of M1; distance between R3 and R4 at least twice as long as distance between R4 and R5 | A. (R.) nothofagetorum Alexander |
16’ | Length of basal deflection of M1 longer than M1+2; distance between R3 and R4 shorter than 1.8× distance between R4 and R5 | A. (R.) caleuchus sp. nov. |
17 | Basal deflection of M1 as long as M1+2; marking at end of R1 small (Fig. |
A. (R.) deceptus sp. nov. |
17’ | Length of basal deflection of M1 shorter than M1+2; marking at end of R1 large (Fig. |
A. (R.) stigmaticus sp. nov. |
18 |
Branches of clasper of gonostylus of different lengths (Fig. |
A. (R.) nonnullus Alexander |
18’ |
Branches of clasper of gonostylus subequal in length (Fig. |
19 |
19 |
19 Apices of crossveins m-cu and r-m desclerotized (Fig. |
A. (R.) apiculatus Alexander |
19’ |
Cross-veins m-cu and r-m uniformly sclerotized (Fig. |
20 |
20 |
Cross-vein m-cu inserted in M3+4; lobule longer than stem of lobe of gonostylus (Fig. |
A. (R.) falcatus sp. nov. |
20’ |
Cross-vein m-cu inserted in bM or at fork; lobule half-length of stem of lobe of gonostylus (Fig. |
21 |
21 |
Knob of halter narrow (Fig. |
22 |
21’ |
Knob of halter wide (Fig. |
23 |
22 |
Tips of mesal lobes of gonocoxite short (Fig. |
A. (R.) billinghami sp. nov. |
22’ |
Tips of mesal lobes of gonocoxite long (Fig. |
A. (R.) theischingeri sp. nov. |
23 |
Female cercus subequal to or shorter than hypogynial valves (Fig. |
A. (R.) immaculatus sp. nov. |
23’ |
Female cercus longer than hypogynial valves (Fig. |
A. (R.) nullus Alexander |
A proper understanding of the phylogenetic relationships within the subgenus A. (Ramphoneurus) depends on a broad taxonomic sampling including representatives of all subgenera of Amphineurus. With that in mind, we included in the analysis representative species of all the subgenera of Amphineurus from the Neotropical and Australasian regions (26 species in total).
Here the characters and their states are defined (the number of steps of each character as they were optimized in the chosen tree is shown within square brackets). In some cases, the understanding of the morphological features under description requires examination of the accompanying illustrations. The data matrix, with a total of 54 terminal taxa and 112 characters, is shown in Supplementary Material 1.
0 Size of rostrum: (0) shorter than remainder of the head; (1) longer or equal to remainder of the head. [1]
1 Shape of rostrum: (0) bifurcated (Fig.
2 Shape of bifurcation: (0) short appendices, shorter than half-length of labrum (Fig.
3 Size of labrum: (0) longer than half the size of the hypopharynx (Fig.
4 Setae of antennae: (0) few setae (Fig.
5 Width of scape: (0) wider or equal to pedicel; (1) thinner than pedicel. [4]
6 Width of terminal flagellomere: (0) as wide as penultimate flagellomere; (1) 1.5× wider than the penultimate flagellomere. [2]
7 Long setae on anterior region of prescutum: (0) absent; (1) present (Fig.
8 Two rows of short setae on dorsal prescutum: (0) absent; (1) present (Fig.
9 Width of prescutum (including presutural scutum): (0) as wide as postsutural scutum; (1) shorter than postsutural scutum. [2]
10 Central dark line on dorsal scutum: (0) absent; (1) present. [3]
11 Lateral dark lines on dorsal scutum: (0) absent; (1) present. [4]
12 Protuberance on scutum: (0) absent; (1) present (Fig.
13 Shape of scutellum in lateral view: (0) with a notch (Fig.
14 Long setae on anteropronotum: (0) absent; (1) present (Fig.
15 Long setae on anepimeron: (0) absent; (1) present (Fig.
16 Height of anepisternum: (0) less than katepisternum (Fig.
17 Notch on margin of anepisternum: (0) present (Fig.
18 Anterodorsal margin of anepisternum: (0) at level or below posterodorsal margin (Fig.
19 Shape of posterior basalare sclerite: (0) less than 2× longer than wide; (1) more than 3× longer than wide. [1]
20 Laterotergite: (0) divided; (1) undivided. [1]
21 Long setae on mediotergite: (0) absent; (1) present (Fig.
22 Fissure in dorsal mediotergite: (0) absent; (1) present (Fig.
23 Midcoxa: without membranous opening (Fig.
24 Knob of halter: (0) less than 2× longer than wide (Fig.
25 Dense vestiture on wing: (0) absent; (1) present. [1]
26 Flattened scales on wing: (0) absent; (1) present. [1]
27 Dense concentration of setae on distal region of wing: (0) present (Fig.
28 Markings at tip of wing: (0) absent (Fig.
29 Marking near apex of vein R1: (0) absent; (1) present (Fig.
30 Markings at forks of radial veins: (0) absent; (1) present (Fig.
31 Pterostigma (marking near vein R2): (0) absent; (1) present (Fig.
32 Marking at vein Rs: (0) absent; (1) present (Fig.
33 Marking at crossvein m-cu: (0) absent; (1) present (Fig.
34 Marking on anal vein: (0) absent; (1) present (Fig.
35 Tip of distal region of vein R5: (0) darker than other veins (Fig.
36 Tip of vein R1: (0) complete (Fig.
37 Base of vein Rs: (0) without spur; (1) with long spur. [1]
38 Tip of crossvein r-m: (0) complete (Fig.
39 Vein M1+2: (0) complete (Fig.
40 Tip of vein bM: (0) complete (Fig.
41 Tip of crossvein m-cu: (0) complete (Fig.
42 Bifurcation of vein Rs: (0) R2+3+4 and R5 (Fig.
43 Vein R2: (0) desclerotized (Fig.
44 Length of vein R2+3: (0) less than 2× longer than R2+3+4; (1) at least 2× longer than R2+3+4. [1]
45 Length of crossvein r-m: (0) longer than basal deflection of R5; (1) shorter or equal to basal deflection of R5. [5]
46 Cell d: (0) closed; (1) open. [4]
47 Length of discal cell: (0) at least 2× longer than wide; (1) less than 2× longer than wide. [2]
48 Vein M3+4: (0) bifurcated; (1) non-bifurcated. [2]
49 Shape of crossvein m-cu: (0) straight (Fig.
50 Position of crossvein m-cu: (0) inserted more distal than fork of Rs; (1) inserted at level of fork of Rs; (2) inserted more proximal than fork of Rs. [6]
51 Shape of anal vein: (0) straight (Fig.
52 Cercus: (0) without membranous projection; (1) with membranous projection. [1]
53 Shape of cercus: (0) slightly curved dorsad (Fig.
54 Shape of male tergite IX: (0) wider than long; (1) longer than wide. [2]
55 Distal half of male tergite IX: (0) dark (Fig.
56 Sclerotized projection at apex of male tergite IX anchor-shaped: (0) absent; (1) present (Fig.
57 Caudal projection in dorsal margin of male tergite IX: (0) absent; (1) present. [1]
58 Number of humps in dorsal margin of male tergite IX: (0) 0; (1) 2; (2) 3; (3) 4. [8]
59 Height of two medial humps in dorsal margin of male tergite IX: (0) tall; (1) short. [3]
60 Height of two lateral humps in dorsal margin of male tergite IX: (0) tall; (1) short. [1]
61 Shape of median notch of male tergite IX: (0) V-shaped (Fig.
62 Size of median notch of male tergite IX: (0) at least 2× wider than deep (Fig.
63 Foramens in tergite X: (0) absent; (1) present. [1]
64 Gonostylus: (0) non-bifurcated; (1) bifurcated in lobe and clasper. [1]
65 Symmetry of claspers of gonostylus: (0) symmetrical; (1) asymmetrical. [1]
66 Base of clasper of gonostylus: (0) pedunculated; (1) wide. [1]
67 Basal spur in clasper of gonostylus: (0) absent; (1) present. [1]
68 Lateroapical spur in clasper of gonostylus: (0) absent; (1) present. [1]
69 Dorsoapical spur in clasper of gonostylus: (0) absent; (1) present. [1]
70 Dense concentration of setae in clasper of gonostylus: (0) absent; (1) present (Fig.
71 Clasper of gonostylus: (0) non-bifurcated; (1) bifid. [1]
72 Position of fork of clasper of gonostylus: (0) very proximal; (1) in more distal half. [1]
73 Shape of medial branch of clasper of gonostylus: (0) straight (Fig.
74 Size of branches of clasper of gonostylus: (0) branches of same length (Fig.
75 Size of curved branch of clasper of gonostylus: (0) not surpassing straight branch (Fig.
76 Shape of lateral branch of clasper of gonostylus: (0) straight (Fig.
77 Lobe of gonostylus: (0) non-bifurcated; (1) bifid. [2]
78 Position of fork of lobe of gonostylus: (0) very proximal; (1) in more distal half. [1]
79 Protuberance in stem of lobe of gonostylus: (0) absent; (1) present (Fig.
80 Length of lobule: (0) shorter than half-length of stem of lobe of gonostylus (Fig.
81 Width of lobule: (0) wider than stem of lobe of gonostylus (Fig.
82 Shape of lobule: (0) longer but less than 4× longer than wide (Fig.
83 Tip of lobule: (0) rounded (Fig.
84 Gonocoxite: (0) non-bifurcated; (1) bifid. [1]
85 Mesal lobe: (0) absent; (1) present. [2]
86 Height of mesal lobe: (0) tall; (1) short. [1]
87 Color of mesal lobe: (0) unpigmented (Fig.
88 Setae in mesal lobe: (0) lateral rows (Fig.
89 Lateral non-sclerotized bulge in mesal lobe: (0) absent; (1) present (Fig.
90 Sclerotized tip of mesal lobe: (0) absent; (1) present (Fig.
91 Shape of sclerotized tip of left mesal lobe: (0) straight (Fig.
92 Shape of sclerotized tip of right mesal lobe: (0) straight (Fig.
93 Base of right mesal lobe: (0) asymmetrical (Fig.
94 Lateral process of sheath of aedeagus: (0) well-developed; (1) absent. [1]
95 Protuberance in sheath of aedeagus: (0) absent; (1) present (Fig.
96 Position of lateral process of sheath of aedeagus: (0) fused laterally with aedeagus; (1) detached from aedeagus. [1]
97 Coloration on detached sheath of aedeagus: (0) absent (Fig.
98 Size of coloration on detached sheath of aedeagus: (0) only at apex (Fig.
99 Setae on detached sheath of aedeagus: (0) absent; (1) present (Fig.
100 Shape of detached sheath of aedeagus: (0) straight; (1) angulated (Fig.
101 Shape of angulated sheath of aedeagus: (0) one-folded (Fig.
102 Spur of sheath of aedeagus: (0) absent; (1) present (Fig.
103 Size of spur of sheath of aedeagus: (0) short (Fig.
104 Shape of spur of sheath of aedeagus: (0) triangular (Fig.
105 Height of spur of sheath of aedeagus: (0) short (Fig.
106 Lateral branches in sheath of aedeagus: (0) absent; (1) present. [2]
107 Shape of lateral branches: (0) straight; (1) curved. [1]
108 Length of lateral branches: (0) not surpassing aedeagus; (1) surpassing aedeagus. [1]
109 Orientation of lateral branches: (0) divergent and surpassing axis of sheath of aedeagus; (1) vertical and not surpassing axis of sheath of aedeagus. [1]
110 Tips of lateral branches: (0) unpigmented; (1) dark. [1]
111 Posterior lateral branches on sheath of aedeagus: (0) absent; (1) present. [2]
The parsimony analysis with equal weights resulted in one most parsimonious tree of 209 steps, Consistency Index (CI) = 0.58 and Retention Index (RI) = 0.86. The analyses with implied weighting (k ranging 1–6) resulted in only one most parsimonious tree with the same topology, shown in Figs
Most parsimonious tree (part) yielded by parsimony analysis with implied weighting (K=3), depicting recovered relationships among outgroup taxa. Numbers at tree branches designate nodes (see 4.2.). — Abbreviations: Aus, Australia; NC, New Caledonia; NG, New Guinea; NZ, New Zealand, SAm, South America; Tas, Tasmania.
To discuss the distribution of the characters in the tree, we use the result of the analysis with implied weighting (with k=3), focusing on the more relevant characters. We believe that our extended outgroup sampling has an enormous heuristic value for future research on the genus Amphineurus, and although we will not discuss here the synapomorphies of the clades recovered in the outgroup, we have numbered all the nodes of the cladogram for future reference. In the lines below, we discuss the synapomorphies and the relationships recovered for the subgenus A. (Rhamphoneurus) (Node 25, Fig.
The synapomorphies of each clade (as provided by the analysis with implied weighting with k=3) are listed within square brackets in the discussion of each node in a standardized way: [number of character: plesiomorphic state – synapomorphic state]. Homoplastic apomorphic states are indicated by an asterisk (*).
Node 25 — subgenus A. (Rhamphoneurus). Synapomorphies: [50: 2-1*]; [77: 0-1*]; [78: 0-1].
The clade is supported by the bifurcated lobe of gonostylus [77: 1]. This feature is present also in clade 9 (outgroup), but there are differences. In A. (Rhamphoneurus), the bifurcation of the lobe is a lobule [78: 1]. Besides, the position of crossvein m-cu [50: 1] reinforces this clade.
In our analysis, the species A. extraordinarius is the sister group of all the other species of A. (Rhamphoneurus).
Amphineurus extraordinarius has M3+4 bifurcated, as in the Neotropical A. castreoensis. However, this resemblance in the medial sector is a symplesiomorphy. A considerable difference between them is the open discal cell of A. extraordinarius as opposed to a closed one in A. castroensis. The very short M3+4 of A. extraordinarius is a necessary step to the probable capture of M3 in Node 26. In our view, this species is a representative of a short-rostrum lineage of A. (Rhamphoneurus), and we expand the diagnosis of the subgenus to include it.
Node 26. Synapomorphies: [0: 0-1]; [1: 1-0*]; [5: 0-1*]; [48: 0-1*]; [55: 1-0*]; [87: 0-1*]; [88: 2-0*]; [98: 1-0*]; [100: 0-2*].
The subgenus A. (Rhamphoneurus) was erected by
The terminalia in this clade is very different from that of other Amphineurus. The distal half of the male tergite IX is dark [55: 0] (Fig.
Node 27. Synapomorphies: [31: 1-0*]; [37: 1-0]; [45: 1-0]; [73: 1-0*].
This clade is supported by wing features such as crossvein r-m longer than the basal deflection of R5 [45: 0] and vein R2 faded [37: 0] (Fig.
Node 28. Synapomorphies: [9: 0-1*]; [11: 0-1*]; [91: 0-1*].
This clade groups the species A. (R.) nullus and A. (R.) falcatus sp. nov. The main synapomorphy is the asymmetrical sclerotized tips of the mesal lobes [91: 1]. Asymmetrical structures are not uncommon in Amphineurus. These species show one tip of the mesal lobe straight and the other bent (Fig.
Node 29. Synapomorphies: [10: 1-0*]; [50: 1-0*].
This clade is supported by a simple scutum, without dorsal markings [10: 0]. Also, these species do not show other thoracic markings, an uncommon condition for this subgenus. This clade has species with translucent wings with great concentration of setae (Fig.
Node 30. Synapomorphies: [3: 0-1]; [14: 1-0*]; [18: 0-1]; [19: 0-1]; [24: 0-1]; [31: 0-1*]; [35: 0-1*].
Amphineurus (R.) theischingeri sp. nov. and A. (A.) billinghami sp. nov. are species with high morphological similarity. These species have very similar wings and male terminalia, although the length of the mesal lobes is crucially different. The node is supported by the labrum shorter or equal to half the size of the hypopharynx [3: 1] (Fig.
Other synapomorphies are seen in the thorax: notably long setae on the anteropronotum [14: 0], the anterodorsal margin of the anepisternum above the posterodorsal margin [18: 1], and the posterior basalare sclerite more than 3× longer than wide [19: 1] (Fig.
Node 31. Synapomorphies: [11: 0-1*]; [36: 0-1*]; [38: 0-1]; [39: 0-1]; [41: 0-1]; [74: 0-1*].
This clade is defined by the presence of faded veins in the middle of the wing (Fig.
Despite the co-occurrence, these faded sections are independent events in unrelated veins such as the tip of vein R1 [36: 1], vein M1+2 [39: 1], the insertion of m-cu [41: 1], and the distal section of r-m [38: 1]. Other evidence to the independence of these evolutive novelties is the reversion of some of them independently. The desclerotization of the tip of R1 is reversed in Node 44. These transformations are common in the subgenus. In Node 27, independently, there are species with vein R2 faded.
In addition, this clade is characterized by a distinct clasper of gonostylus (Fig.
Node 32. Synapomorphies: [13: 0-1]; [17: 0-1*].
In the description of A. (R.) apiculatus,
Node 33. Synapomorphies: [4: 0-1*]; [61: 0-1*]; [82: 0-2*]; [98: 0-1*]; [100: 2-1*].
This clade represents the major radiation of the subgenus. The main synapomorphy is the angulated sheath of aedeagus [100: 1]. The shape of the sheath (Fig.
The male terminalia is also characterized by the U-shaped tergite IX [61: 1] (Fig.
Node 34. Synapomorphies: [29: 0-1]; [30: 0-1]; [32: 0-1*]; [33: 0-1*]; [34: 0-1*]; [62: 0-1]; [80: 0-1*]; [88: 0-2*]; [90: 0-1]; [101: 0-1].
This clade is supported by the presence of dark markings on the wing (Fig.
The clade is also sustained by synapomorphies in the male terminalia: the curved shape of median notch of tergite IX [62: 1] and the long lobule [80: 1] (Fig.
Node 35. Synapomorphies: [5: 1-0*]; [12: 0-1*]; [14: 1-0*]; [15: 0-1*]; [28: 0-2]; [49: 0-1]; [51: 0-1*].
This clade is defined by the tip of the wing with markings around the radial veins [28: 2]. The three species of this clade have crossvein m-cu angulated [49: 1]. The strongly sinuous anal vein [51: 1] is a synapomorphy of this clade (Fig.
Another synapomorphy is the scutum with a protuberance [12: 1] (Fig.
Node 36. Synapomorphies: [16: 0-1]; [22: 0-1]; [44: 0-1]; [45: 1-0*]; [50: 1-2*].
This clade groups together A. (R.) glabristylatus and A. (R.) chiloeanus. These two species have different sheaths of the aedeagus, but very similar wings. Both wings have vein R2+3 at least 2× longer than R2+3+4 [44: 1], crossvein r-m shorter than the basal deflection of R5 [45: 0] and crossvein m-cu inserted more proximal than the fork of Rs [50: 2]. Besides wing features, the clade is supported by the anepisternum higher than the katepisternum [16: 1] and by the presence of a fissure in the dorsal mediotergite [22: 1] (Fig.
In the description of A. (R.) chiloeanus,
Node 37. Synapomorphies: [27: 0-1]; [28: 0-1]; [55: 0-1*]; [70: 0-1]; [76: 0-2]; [81: 0-1*]; [89: 0-1].
All specimens of the species of this clade were originally identified as A. (R.) nothofagetorum by C.P. Alexander. However, detailed morphological study uncovered a higher diversity, and three new species are placed in this clade along with A. (R.) nothofagetorum. Node 37 groups the species with darkened distal quarters of the wings [28: 1] (Fig.
Another important synapomorphy is the dorsal margin of the male tergite IX without a dark region [55: 1]. The tergite (Fig.
Node 38. Synapomorphies: [21: 0-1*]; [40: 0-1].
This clade is supported by faded veins near the fork of bM [40: 1] (Fig.
Node 39. Synapomorphies: [5: 1-0*]; [53: 0-1*]; [75: 0-1].
This node groups A. (R.) stigmaticus sp. nov. and A. (R.) caleuchus sp. nov. The main synapomorphy is the long medial branch of the clasper of gonostylus [75: 1] (Fig.
Node 40. Synapomorphies: [7: 0-1*]; [11: 1-0*]; [95: 0-1].
This node represents a major branch of the subgenus. The defining synapomorphy of this node is a protuberance in the sheath of aedeagus [95: 1] (Fig.
Node 41. Synapomorphies: [14: 1-0*]; [82: 2-1*]; [102: 0-1].
This clade is defined by a distinctive spur in the sheath of aedeagus [102: 1]. This distal spur of the sheath of aedeagus (Fig.
Node 42. Synapomorphies: [54: 0-1]; [56: 0-1]; [76: 0-1]; [79: 0-1]; [80: 0-1*]; [83: 0-1]; [85: 1-0*]; [99: 0-1]; [103: 0-1].
This clade groups A. (R.) sanus and A. (R.) anchoralis sp. nov. These two species are very similar to each other and share many apomorphies. The most distinguishable synapomorphy is the anchor-shaped sclerotized projection on male tergite IX [56: 1] (Fig.
The detached sheath of aedeagus of these species is peculiar (Fig.
Node 43. Synapomorphies: [4: 1-0*]; [53: 0-1*]; [73: 1-0*]; [88: 0-1*]; [91: 0-1*]; [92: 0-1*].
This clade shows great morphological variability, especially in the male terminalia. Of the seven species in this group, however, only A. (R.) insanus has been previously described.
The main characteristic of this clade is the strong asymmetry of the mesal lobes. In this clade, the left mesal lobe has a folded tip [91: 1], while the right one is curved [92: 1] (Fig.
The asymmetry in the male terminalia is not uncommon in Amphineurus (e.g., A. (Nesormosia)) nor in the related genus, Maietta (
In addition, the medial branch of the clasper of gonostylus is straight [73: 0]. This feature is observable also in Node 27 (Fig.
Node 44. Synapomorphies: [2: 0-1]; [14: 0-1*]; [23: 0-1]; [36: 1-0*]; [105: 0-1].
This clade is represented by species with a tall spur on the sheath of aedeagus [105: 1] (Fig.
Node 45. Synapomorphies: [6: 0-1*]; [21: 0-1*].
This node groups A. (R.) podenasi sp. nov. and A. (R.) oosterbroeki sp. nov. These species are notable for the dilated last flagellomere [6: 1] (Fig.
Node 46. Synapomorphy: [81: 0-1*].
This node is supported by the lobule thinner than the stem of the lobe of gonostylus [81: 1] (Fig.
Node 47. Synapomorphies: [7: 1-0*]; [104: 0-1].
This node is supported by the quadrangular shape of the spur of the sheath of aedeagus [104: 1] (Fig.
Node 48. Synapomorphy: [93: 0-1].
This node groups A. (R.) insanus and A. (R.) alexanderi sp. nov. These species are so similar that all specimens of A. (R.) alexanderi sp. nov. were misidentified by C.P. Alexander as A. (R.) insanus. The defining synapomorphy of this clade is the right mesal lobe with symmetrical sides in the base [93: 1] (Fig.
The label data of the Amphineurus specimens suggest a univoltine development. The adults are found from the end of November to April, but the majority of species occur from December to January. There is a wide range of tolerated altitude, from sea-level to 1600 m. Most specimens and species were collected in humid environments. Amphineurus (A.) submolophilinus is suggested to be found near streams (
The larvae of A. (A.) hudsoni have been found in wet decayed fallen leaves of Nikau (Rhopalostylis sapida Wendl and Drude), an endemic New Zealand palm (
In South America, data is more scarce. However, A. extraordinarius is associated with trees of Araucaria Kuntze (
The biogeography of the genus Amphineurus is an old topic.
Although the focus of our study is the subgenus Amphineurus (Rhamphoneurus) (Fig.
Concerning taxa distributed in Australasia and South America, there is a recurrent asymmetry, also not expected by vicariance driven by the fragmentation of Gondwana, in which Australasian taxa (notably, New Zealand taxa) constitute a grade (i.e., are paraphyletic) relative to South American taxa. This asymmetry has been found, for instance, in phylogenetic studies of other components of this Trans-Pacific Biota (such as the genus Aphrophila, recently revised by
The asymmetry just mentioned, and other general distributional patterns non-congruent with the sequence of breakup of Gondwana can be interpreted as evidence of a heterogeneous set of ancient, pre-rift distribution ranges in the area. Recent phylogenetic studies including both fossil and recent representatives of Tipuloidea (Lukaschevich and Ribeiro 2019) indicate that the main Tipuloidea lineages (family level clades) have differentiated, at least, from the Triassic (stem lineages) to Upper Jurassic times (more derived families). During this time interval, a lot of pre-rift differentiation must have occurred before the final continental divisions. The separation of continental landmasses is only the final result of long-lasting tectonic events, which caused a lot of in situ differentiation by altering the local landscapes.
One alternative explanation could be by a posteriori dispersal from Australasia to South America, but the poor dispersal capabilities of some (if not all) of the groups in question and their very particular habitat needs (for instance, larvae of Aphrophila craneflies are restricted to fast-flowing waters in mountainous areas) does not support such views. In addition, assuming independent trans-oceanic dispersal seems odd with a unifying view of biogeography as the result of an evolving geological setting (
In the words of
We are very grateful to Dr. Wayne Mathis, for the loan of the specimens used in this study (as well as for his unconditional support to the work of GCR for many years) when he was still working as curator of Diptera at the NMNH, Washington. We also thank one anonymous reviewer, Dr. Levente-Péter Kolcsár, and subject editor, Dr. Bradley Sinclair, for detailed reviews and corrections that greatly contributed to the final version of this paper. We thank Daniel Dias Dornelas do Carmo for assistance in the Bayesian analysis. This study was financed by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) – Finance Code 001 (RRS) and by FAPESP (grant 2017/16305-6 to DS; grant 2010/00557-7 and 2020/02844-5 to GCR).
Morphological matrix
Data type: .ss
Explanation note: The file contains the matrix used in the phylogenetic analysis.